From the Atlantic Forest of Serra De Paranapiacaba, Southestern Brazil

ARTICLE The Muscidae (Diptera) from the Atlantic Forest of Serra de Paranapiacaba, southestern Brazil

Silvio Shigueo Nihei¹³; Anderson Tatsuki Tamakoshi¹⁴; Priscylla Moll¹⁵ & Rodrigo Ferreira Krüger²⁶

¹ Universidade de São Paulo (USP), Instituto de Biociências (IB‑USP), Departamento de Zoologia. São Paulo, SP, Brasil. ² Universidade Federal de Pelotas (UFPEL), Instituto de Biologia (IB), Departamento de Microbiologia e Parasitologia (DEMP). Pelotas, RS, Brasil. ³ ORCID: http://orcid.org/0000-0002-6887-6282. E‑mail: [email protected] ⁴ ORCID: http://orcid.org/0000-0003-3954-1018. E‑mail: [email protected] ⁵ ORCID: http://orcid.org/0000-0002-0226-7732. E‑mail: [email protected] ⁶ ORCID: http://orcid.org/0000-0003-1040-8299. E‑mail: [email protected]

Abstract. A first survey of Muscidae in the State of São Paulo (Southeastern Brazil) is presented here with a one-year of Malaise trap collecting from August 2010 to July 2011 at the Biological Reserve Alto da Serra de Paranapiacaba (23°46 00 ‑23°47 10 S, ′ ″ ′ ″ 46°18 20 ‑46°20 40 W, 750‑891 m of altitude). A total of 1,284 individuals of muscids were collected, and 15 genera and 39 ′ ″ ′ ″ species of Muscidae were identified. So far, only one muscid species had been recorded to the Reserve, which now has its Muscidae diversity increased to 40 species. Thirteen species are new records for the State of São Paulo. With this, the number of species of Muscidae species known to occur in the State of São Paulo is increased to 169. The interval between November and February was higher in number of individuals and number of species. Muscidae presented a seasonal pattern, with more abundance and diversity in that interval. The study area is covered by secondary forest and very close to São Paulo metropolitan area, and the composition of the fauna of Muscidae signalizes this environment changing and anthropic stress with nine species with synanthropic habits, two of them are typically synanthropic species.

Keywords. Diversity; Inventory; Neotropical region; Seasonality; State of São Paulo; Survey.

INTRODUCTION the different Brazilian biomes and states. For example, while there is a voluminous literature on The family Muscidae is worldwide distribut- Muscidae for the States of Paraná and Rio de ed and include about 5,200 described species in Janeiro (e.g., Carvalho et al., 1984; D’Almeida, 187 genera (Pape et al., 2011). In the Neotropical 1991; Pamplona et al., 2000; Oliveira et al., 2002; Region there are over 800 species and 85 gen- Costacurta et al., 2003; Couri & Carvalho, 2005; era (Couri & Carvalho, 2005). The family is greatly Leandro & D’Almeida, 2005; Rodríguez-Fernández abundant in all continents, occurring in most oce- et al., 2006; Couri & Barros, 2009), the most rich anic islands, and also in the Arctic and regions of and populous Brazilian State remains with the tundra vegetation (Huckett & Vockeroth, 1987). Muscidae diversity knowledge still incipient. Muscid flies have diverse feeding habits, includ- Furthermore, it is relatively scarce the Muscidae ing predators, phytophagous, saprophagous, surveys in the Neotropics based on the use of anthophilous, parasites and haematophagous non-baited collecting techniques (e.g., Carvalho (Couri, 1999; Savage & Vockeroth, 2010; Carvalho & Couri, 1991; Costacurta et al., 2003; Rodríguez- et al., 2012). Fernández et al., 2006; Krüger et al., 2010; Zafalon- There are a number of studies with invento- Silva et al., 2018). The present study is the first ex- ry of Diptera in the Neotropics (e.g., Carvalho & haustive survey of Muscidae for the State of São Couri, 1991; Couri et al., 2000; Pamplona et al., Paulo based on non-baited collectings. Previous 2000; Krüger et al., 2010; Borkent et al., 2018; surveys focused on urban and/or cadaveric fauna Zafalon-Silva et al., 2018), on the other hand, sev- (Linhares, 1981; Dias et al., 2014; Cavallari et al., eral studies focused on groups with agricultural 2015). In Brazil, 358 species are described and or forensic importance (e.g., Ferraz et al., 2009; recorded, 156 of these occur in the State of São Mulieri et al., 2011; Garcia et al., 2003; Cavallari Paulo (Lowenberg-Neto & Carvalho, 2013, with et al., 2015; Carvalho et al., 2017; Olea et al., 2017). updates from Pereira-Colavite & Carvalho, 2012; With regard to Muscidae, there are a number of Cavallari et al., 2015; Fogaça & Carvalho, 2018). inventories in Brazil, but unbalanced between In this article we surveyed the Muscidae of the

Pap. Avulsos Zool., 2021; v.61: e20216168 ISSN On-Line: 1807-0205 http://doi.org/10.11606/1807-0205/2021.61.68 ISSN Printed: 0031-1049 http://www.revistas.usp.br/paz ISNI: 0000-0004-0384-1825 http://www.scielo.br/paz Edited by: Carlos José Einicker Lamas Received: 07/05/2021 Accepted: 26/05/2021 Published: 30/07/2021 Pap. Avulsos Zool., 2021; v.61: e20216168 Nihei, S.S. et al.: Muscidae from Paranapiacaba 2/12

Biological Reserve Alto da Serra de Paranapiacaba (State MATERIAL AND METHODS of São Paulo, southeastern Brazil) in the Atlantic Forest biome, and analyzed its abundance, diversity and sea- Study area and collection sonality throughout a one-year collecting period. So far, only one muscid species had been recorded to the This survey took place in the Biological Reserve Alto Biological Reserve of Paranapiacaba (Townsend, 1927; da Serra de Paranapiacaba (Fig. 1) (23°46 00 , 23°47 10 S; ′ ″ ′ ″ Lowenberg-Neto & Carvalho, 2013). 46°18 20 , 46°20 40 W, 750‑891 m of altitude), located ′ ″ ′ ″

Figure 1. Location of the Biological Reserve Alto da Serra de Paranapiacaba, State of São Paulo, southeastern Brazil (23°46 00 , 23°47 10 S; 46°18 20 , 46°20 40 W, ′ ″ ′ ″ ′ ″ ′ ″ 750‑891 m of altitude). Nihei, S.S. et al.: Muscidae from Paranapiacaba Pap. Avulsos Zool., 2021; v.61: e20216168 3/12 in the municipality of Santo André, State of São Paulo, tervals (95%) as a function of abundance was performed Brazil. The reserve is situated slightly under the Capricorn in R using the statistical package iNEXT (Hsieh et al., Tropic (23°27 S), in a transition area between the Tropical 2016) based on the methodologies proposed by Colwell ′ and the Subtropical climates zones (Gutjahr & Tavares, et al. (2012) and Chao et al. (2014). 2009). According to Köppen’s classification, the climate Collector curves were obtained for the Biological type is Cwa, temperate and humid, with dry winter Reserve Alto da Serra de Paranapiacaba and for each sea- and hot summer. The mean temperature at summer sonal season within the same area. (December-March) is 20 , and 14 on winter (June- ℃ ℃ September), with temperature range of 6 (Gutjahr & ℃ Tavares, 2009). The area is predominantly covered with RESULTS secondary Atlantic Forest under different successional stages, and that altered flora is due to historical anthrop- Within the one-year of Malaise trap collected ma- ic intervention (Sugiyama et al., 2009), as this area is very terial, seven families of Calyptrata were surveyed: close to São Paulo metropolitan area. Calliphoridae, Fanniidae, Mesembrinellidae, Muscidae, Insects were collected using Malaise traps, installed Rhinophoridae, Sarcophagidae and Tachinidae. Those at three different point of the Reserve: trap 1 was locat- seven families accounted a total of 3,451 individuals, ed at 23°46 38 S, 46°18 42,7 W; trap 2 at 23°46 44.6 S, with Muscidae as the most abundant, with 1,284 indi- ′ ″ ′ ″ ′ ″ 46°18 40,2 W; and trap 3 at 23°46 45,7 S, 46°18 29,2 W. viduals (37%). Excluding the genera formerly belonging ′ ″ ′ ″ ′ ″ Traps were installed and material had been collected to “Coenosiinae” (sensu Carvalho et al., 2005), the collect- continuously from August 2010 to July 2011. Sampling ed material of Muscidae was restricted to 397 individu- vessels were replaced monthly, and the collected materi- als, and a total of 15 genera and 39 species of Muscidae al taken to the lab for sorting and identification. were identified (Table 1 and Appendix) from the three subfamilies of Muscidae: Muscinae, Cyrtoneurininae and Mydaeinae (following the subfamilial classification sensu Identification Haseyama et al., 2015). Thirteen species are new records for the State of São Paulo: Dolichophaonia machadoi The specimens were recognized until the genera and (Albuquerque, 1958), D. simplex (Albuquerque, 1958), species level, using identification keys in Carvalho (2002). Helina discreta (Wulp, 1896), H. gigantea Albuquerque, When necessary, taxonomic revisions were consulted, 1956, H. regobarrosi Albuquerque, 1958, Neomuscina containing updated identification keys and descriptions anajeensis Pereira-Colavite & Carvalho, 2012, N. trans- (e.g., Pereira-Colavite & Carvalho, 2012; Nihei & Carvalho, porta Snyder, 1949, Neurotrixa sulina Costacurta & 2007; among others). All studied material was pinned Carvalho, 2005, Phaonia advena Snyder, 1957, P. annu- and, in most cases, males and/or females were dissect- lata (Albuquerque, 1957), P. grandis (Couri, 1982), P. len- ed to study their terminalia, which were compared to tiginosa Snyder, 1957 and Polietina minor (Albuquerque, drawings and descriptions in the literature. The dissect- 1956). With this, the number of species of Muscidae ed terminalia were cleared with 10% KOH solution, then known to occur in the State of São Paulo is increased to neutralized with acetic acid solution and washed with 169. water; after examination, they were stored in microvials Quantitative data analysis reveals that the interval with glycerine and pinned with the respective specimen. between November and February had both more indi- The material was deposited at the Museu de Zoologia, viduals (Fig. 2) and more species (Fig. 3) collected. Those Universidade de São Paulo, São Paulo (MZSP). three-month period have a sequence of 60‑55‑56 individuals, while other months reached up to 36 individuals. For species number, we had a variation of 14 to 18 spe- Data analyses cies. Shannon diversity index confirms summer with the highest diversity, followed by spring, autumn and winter Quantitative data were analyzed by calculating (Table 2). On the other hand, Berger-Parker dominance Shannon diversity and Berger-Parker dominance indexes shows summer with the lowest rate, followed by spring (Magurran, 1988) to compare the diversity and propor- and autumn, and with winter with the highest rate. In tional abundance of species among the four stations of the Berger-Parker dominance, the lower the index the higher year. These calculations were performed in the software the dominance of such taxon (Table 2). DivEs v2.0 (Rodrigues, 2005). To analyze the seasonality The estimated values of Chao for the richness of mus- abundance in the four seasons of the year, we conducted cid species in the Biological Reserve was 65.009 ± 12.198. the Kruskal-Wallis statistical test for non-parametric sam- The Chao estimator is accurate in its lower limit and based ples using the software Past version 2.17c (Hammer et al., on it, we can infer that about 52 species are expected in 2001). We have applied a Kruskal-Wallis post-test called the area, seven more than the observed richness. This Bonferroni Correction, used to reduce the chances of ob- value can reach 105 species, which is an overestimate taining false-positive results (type I errors) when several due to the 14 singletons (species with only one individu- pairwise tests are made for one dataset (Nakagawa, 2004). al) (31.11%) (Table 3). There is also no trend towards sta- The estimation of the number of species and extrapo- bilization of the collector curve (Fig. 4a), corroborating lation of this variable and their respective confidence in- Chao’s analysis for the Biological Reserve. Pap. Avulsos Zool., 2021; v.61: e20216168 Nihei, S.S. et al.: Muscidae from Paranapiacaba 4/12

Table 1. Species of Muscidae (excluding “Coenosiinae”) from Biological Reserve Alto da Serra de Paranapiacaba with the number of individuals per season in the period August 2010 to July 2011. Asterisks indicate new records to State of São Paulo.

Species Winter Spring Summer Autumn Species Winter Spring Summer Autumn Cyrtoneurina alifusca 1 1 6 1 Neurotrixa sulina* 0 0 1 0 Cyrtoneurina costalis 0 0 0 1 Phaonia advena* 5 16 20 12 Cyrtoneuropsis maculipennis 7 11 41 18 Phaonia annulata* 0 0 1 1 Cyrtoneuropsis polystigma 3 14 21 0 Phaonia grajauensis 0 0 2 0 Dolichophaonia machadoi* 5 6 3 1 Phaonia grandis* 29 25 32 25 Dolichophaonia simplex* 0 1 0 0 Phaonia lentiginosa* 2 5 7 3 Helina discreta* 0 0 3 1 Phaonia shannoni 2 1 1 0 Helina gigantea* 0 2 0 0 Phaonia similata 0 1 0 0 Helina praecipua 1 0 0 0 Phaonia trispila 1 0 0 0 Helina regobarrosi* 0 0 3 0 Philornis amazonensis 0 1 1 0 Morellia humeralis 0 0 1 0 Polietina minor* 0 0 1 0 Morellia nigricosta 0 0 3 0 Polietina prima 0 0 3 3 Mydaea plaumanni 3 1 3 1 Myospila meditabunda 0 1 1 0 Polietina orbitalis 0 0 1 0 Neomuscina anajeensis* 0 1 0 0 Polietina steini 0 0 2 0 Neomuscina atincticosta 0 0 1 0 Polietina univittata 0 0 3 1 Neomuscina inflexa 0 0 2 0 Pseudoptilolepis fulvapoda 1 6 4 1 Neomuscina neosimilis 0 0 2 0 Pseudoptilolepis nudapleura 0 0 1 0 Neomuscina schadei 0 0 0 1 Psilochaeta pampiana 0 0 2 0 Neomuscina transporta* 0 1 0 0 Synthesiomyia nudiseta 0 0 0 1

Table 2. Shannon diversity and Berger-Parker dominance indexes. The high‑ bilization in the collector curve for summer, spring and est rates are highlighted in bold. autumn, while in the winter the collections were suffi- cient (Fig. 4b). Shannon’s index Berger-Parker The seasonality analysis using Kruskal-Wallis sta- Autumn 0,8181 0,3521 tistical test (Table 4) showed a seasonal pattern for the Winter 0,7643 0,5 abundance of Muscidae throughout the year, with sig- Spring 0,9609 0,266 nificative difference between the number of individuals Summer 1,0979 0,2398 distributed in the stations. The difference among the stations found at Kruskal-Wallis test was due to the abun- Table 3. Muscidae richness (Chao) from August 2010 to July 2011 at the dance of individuals in the summer when compared Biological Reserve Alto da Serra de Paranapiacaba (ReBio) in São Paulo, Brazil. to the winter and fall seasons. Furthermore, the circu- Observed values (Obs), estimated (est), standard error of the estimated val‑ lar analysis (Fig. 5) supported the seasonal variation of ues (SE), lower (LL) and upper (UL) limits of the estimated values. Muscidae indicating a clustered distribution (r = 0,913). The five most abundant species (Fig. 6) were Phaonia Richness (Chao) Local/Season grandis (28.17%), Cyrtoneuropsis maculipennis Macquart, Obs est SE LL UL 1843 (19.54%), Phaonia advena (13.45%), Cyrtoneuropsis ReBio 45 65.009 12.198 51.649 105.217 polystigma (Wulp, 1896) (9.13%), and Phaonia lentiginosa Spring 27 37.003 7.942 29.541 66.375 (4,31%). Summer 35 75.230 28.364 46.584 174.720 Autumn 17 27.507 10.125 19.146 68.441 Winter 16 19.948 5.228 16.549 44.380 DISCUSSION

Table 4. Pairwise Kruskal-Wallis analyses. Correction of Bonferroni are in In the Biological Reserve Alto da Serra de gray color. Significative values are underlined. Paranapiacaba, only one muscid species, Philornis mima (Townsend, 1927), had been recorded so far (Townsend, Winter Spring Summer Autumn 1927; Lowenberg-Neto & Carvalho, 2013). With the 39 Winter 0,262 0,0005 0,629 species found in the present survey, the Muscidae diver- Spring 1 0,013 0,532 sity of the Reserve is increased to a total of 40 species. Summer 0,003 0,077 0,0009 Malaise traps are efficient in collecting very active fly- Autumn 1 1 0,006 ing insects that can be intercepted by its tents, especially Hymenoptera and Diptera (Campos et al., 2000). In trop- Estimates of Muscidae richness for each season of ical rain forests of the Neotropical Region, Diptera is the the year, as well as their extrapolation, reveal interesting most collected order when using Malaise traps (Dutra & patterns of sampling effort for this conservation unit. In Marinoni, 1994; Brown, 2005). Costacurta et al. (2003) and Table 3 we can see that there are higher values in the Rodríguez-Fernández et al. (2006) analyzed the composi- summer than in any other season, without a trend of sta- tion, species richness and abundance of Muscidae in six Nihei, S.S. et al.: Muscidae from Paranapiacaba Pap. Avulsos Zool., 2021; v.61: e20216168 5/12 localities in Paraná State, southern Brazil in the Atlantic When considering the main surveys of Muscidae for Forest biome, during one-year with Malaise traps. Krüger southern Brazil (Rodríguez-Fernández et al., 2006; Krüger et al. (2010) surveyed four localities in southern Brazilian et al., 2010; Zafalon-Silva et al., 2018), we observe that grasslands in the Pampa biome using a one-year Malaise the richness in this family, disregarding the Coenosiinae, traps collecting; and Zafalon-Silva et al. (2018) collected is 109 species for Paraná State (Rodríguez-Fernández Muscidae in the 35 areas in the Coastal Plain of the Rio et al., 2006), 59 species for localities in the grasslands Grande do Sul State, southern Brazil. Borkent et al. (2018) and south coast of Lagoa dos Patos (Krüger et al., 2010) surveyed three localities in the Costa Rican cloud forests and 63 species for the coastal plain of the Rio Grande using 13‑month Malaise traps. do Sul (Zafalon-Silva et al., 2018), values that are 31 to

Figure 2. Number of individuals of Muscidae per month for the whole year of study.

Figure 3. Number of species of Muscidae per month for the whole year of study. Pap. Avulsos Zool., 2021; v.61: e20216168 Nihei, S.S. et al.: Muscidae from Paranapiacaba 6/12

132% higher than that found for Biological Reserve of phospecies, the richness in the Biological Reserve of Paranapiacaba. It is worth mentioning that the morphos- Paranapiacaba was of 41 species, values close to those pecies richness values are also much higher than those found by Krüger et al. (2010) and Zafalon-Silva et al. found here, varying from 23% of the fauna in Krüger (2018), 45 and 42 species, respectively. In Paraná, the et al. (2010) to 38% in Rodríguez-Fernández et al. (2006), richness was of 67 species (Rodríguez-Fernández et al., while in this work only four species were not nominally 2006), remaining 63% higher than in Paranapiacaba. identified (8.89%). Therefore, when disregarding mor- The collections carried out at the Biological Reserve of

Figure 4. Species accumulation curve based on the abundance of muscids captured with Malaise traps at the Biological Reserve Alto da Serra de Paranapiacaba, State of São Paulo. The solid line represents the observed richness of the collection and the dashed line the richness estimated by Chao 1. (A) corresponds to the estimates for the Reserve considering all collections. (B) corresponds to the estimates for each season of the year: spring (green), summer (blue), autumn (purple) and winter (red). Nihei, S.S. et al.: Muscidae from Paranapiacaba Pap. Avulsos Zool., 2021; v.61: e20216168 7/12

Figure 5. Circular representation of the number of collected individuals of Muscidae per month, for the whole year of study. Dark grey = rainy season; light grey = dry season. r = concentration index.

Figure 6. Relative frequency of 15 most abundant species of Muscidae sampled. The column “others” comprises 24 low abundant species. Pap. Avulsos Zool., 2021; v.61: e20216168 Nihei, S.S. et al.: Muscidae from Paranapiacaba 8/12

Paranapiacaba used traps that were placed close to each seasonality of Neotropical muscids. Using fruit-baited other (separated by hundreds of meters), and this may traps, Azevedo et al. (2015) found the muscids much less explain the much higher values in Paraná, where the abundant in the rainy season in Barbalha municipality collections were carried out in almost the entire State (State of Ceará, northeastern Brazil), characterized by (separated by hundreds of kilometers). Nevertheless, we Cerrado (savannah) and Caatinga (semi-arid) vegeta- can estimate a great richness for Muscidae in the Atlantic tions. Results of Costacurta et al. (2003) in three localities Forest of São Paulo State, since in a reduced area as the of State of Paraná (southern Brazil) also supports the cor- Biological Reserve of Paranapiacaba, the richness found relation between Muscidae abundance and air humid- was to similar to that found in areas of much larger col- ity: the lower the humidity, the higher the abundance. lections as Rio Grande do Sul, wherein sampled locali- However, as pointed out by Azevedo & Krüger (2013) ties were separated by dozens of kilometers. In fact, the for southern Brazilian calliphorids, each species may re- Chao estimate in Table 3 extrapolates more species to be spond differently (even inversely) to temperature and collected at Paranapiacaba and, in this case, we can con- humidity interactions, therefore, it demands a careful ex- sider estimates higher than the estimated one. Thus, the amination on the species other than all the family. curve of the collector is still increasing for this area. The Paranapiacaba Reserve is covered by second- Interestingly, the fauna found in the Biological ary Atlantic Forest under different successional stages Reserve of Paranapiacaba is more similar to the fauna of (Sugiyama et al., 2009). The composition of the Muscidae Paraná, sharing 31.71% of the species, while only 8% of fauna from the Paranapiacaba Reserve comprises most- Paranapiacaba species were also observed in Rio Grande ly tipically forest species, however there is a number of do Sul and in Paraná. However, none of those shared spe- species with some degree of synanthropic habits. Within cies occurs either in the Coastal Plain of RS or in the Sul- its 39 species, only two species, Myospila meditabunda rio-grandense Shield. And ten species (24.4%) occur in (Fabricius, 1781) and Synthesiomyia nudiseta (Wulp, 1883), all these locations and have a wide distribution in Brazil. are considered tipically synanthropic (Carvalho et al., The seasonal pattern of Muscidae showed the fam- 2005; Lowenberg-Neto & Carvalho, 2013). In Colombia, ily individuals more abundant on summer, based on S. nudiseta was classified as having “strong preference for Kruskal-Wallis statistical test (Table 4) and circular anal- dense human settlements” (Uribe et al., 2010), and in Brazil ysis (Fig. 5). Previous studies have demonstrated that we have it recorded in rural and urban environments and tropical insect abundance decreases in the dry season forest fragments (D’Almeida & Almeida, 1998). Besides, (e.g., Pinheiro et al., 2002; Da Silva et al., 2011), but far less seven species found in Paranapiacaba have some associ- than in temperate zones (Wolda, 1988). The pairwise test ation to human inhabited areas in Brazil: Cyrtoneuropsis comparing summer and spring resulted in a statistically maculipennis, C. polystigma, Morellia humeralis (Stein, relevant difference, but after Bonferroni Correction, this 1919), Mydaea plaumanni Snyder, 1941, Phaonia trispila difference was not considered statistically significant (Bigot, 1885), Pseudoptilolepis fulvapoda Snyder, 1949 (Table 4). On the other hand, Bonferroni tests confirmed and Psilochaeta pampiana (Shannon & Del Ponte, 1926) seasonal pattern of Muscidae with high abundance in (Linhares, 1981; Carvalho et al., 1984; Oliveira, 1986; the summer and less in autumn and winter. Kasai et al., 1990; Lomônaco & Almeida, 1995; Moura Da Silva et al. (2011) and Pinheiro et al. (2002) per- et al., 1997; Pamplona et al., 2000; Carvalho et al., 2002; formed seasonal analyses for insect abundance in the Barbosa et al., 2009). And Cyrtoneuropsis polystigma was Brazilian Cerrado (savannah) and found that most insect also considered symbovine (Oliveira, 1986). However, orders (Coleoptera, Hymenoptera, Hemiptera, Isoptera, some species have contrasting information, for example, etc.) showed a clustered distribution, being more abun- in Colombia (Uribe et al., 2010), C. maculipennis is clas- dant in the wet season. However, in both studies, the or- sified as a species with “complete avoidance of human der Diptera was randomly distributed, contrary to the re- settlements”, as well as Polietina orbitalis (Stein, 1904). sults observed here and focused on Muscidae. This could The occurrence of exotic and/or synanthropic spe- be related to the groups studied in each case, revealing cies in natural preserved areas has been associated to perhaps the most abundant groups of Diptera might edge effect by some authors (Lima-Ribeiro, 2008; Ferraz be equally distributed the whole year, while Muscidae et al., 2010; Barbosa et al., 2014). Furthermore, the com- might be more vulnerable to environmental and/or bi- plex composition of the Muscidae fauna in the Biological otic conditions that affect their abundance. Moreover, Reserve of Paranapiacaba reveals that while this Reserve these studies were performed in different biomes other still bears most of its fauna with typically forest species, than Atlantic Forest. Traditionally, Cerrado is character- the presence of some typically synanthropic species ized by a bimodal rainfall distribution, with two well-de- may signalize that its proximity to São Paulo metropol- fined, dry and wet, seasons (Silva et al., 2008), whereas itan area has imposed an increasing anthropic pressure Atlantic Forest is characterized by having high tem- (Leandro & D’Almeida, 2005). perature and precipitation during the whole year (IBGE, 1992). However, climatic studies in Paranapiacaba have demonstrated that precipitation suffers a decrease in ACKNOWLEDGMENTS December and January (Gutjahr & Tavares, 2009), when Muscidae reached its peaks in both abundance and di- We thank Filipe Gudin for helping with the material versity. Unfortunately, there are just a few studies with sorting; Filipe Gudin, Julia Calhau, Marcos Ribeiro, Mayra Nihei, S.S. et al.: Muscidae from Paranapiacaba Pap. Avulsos Zool., 2021; v.61: e20216168 9/12

Sato, Nelly Araya, Rodrigo Dios and Pedro Souza-Dias Silva, V.C.; Sinclair, B.J.; Skevington, J.H.; Stireman, J.O.; Vilkamaa, P.; for helping in the collecting, and Julia Benetti, Glauco Wheeler, T.; Whitworth, T.; Wong, M.; Wood, D.M.; Woodley, N.; Yau, T; Machado and Daniel Caetano for suggestions on ear- Zavortink, T.J. & Zumbado, M.A. 2018. Remarkable fly (Diptera) diversity lier versions. We also thank the financial support from in a patch of Costa Rican cloud forest: Why inventory is a vital science. FAPESP with research grant (Proc. № 2007/50836‑7 Zootaxa, 4402: 53‑90. DOI for SSN) and scholarships (Proc. № 2007/52314‑7 for Brown, B.V. 2005. Malaise trap catches and the crisis in neotropical ATT and 2010/07185‑8 for PM) and CNPq research Dipterology. American Entomologist, 51(3): 180‑183. DOI grant (Proc. № 562199/2010‑1 for SSN), scholar- Campos, W.G.; Pereira, D.B.S. & Schoereder, J.H. 2000. Comparison of the ship (Proc. № 107159/2011‑2 for ATT) and fellowship efficiency of flight-interception trap models for sampling hymenoptera (Proc. № 309192/2018‑8 for SSN). and other insects. Anais da Sociedade Entomológica do Brasil, 29(3): 381‑389. DOI Carvalho, C.J.B. De. (Ed.). 2002. Muscidae (Diptera) of the Neotropical Region: AUTHORS’ CONTRIBUTIONS Taxonomy. Editora Universidade Federal do Paraná, Curitiba, 287p. Carvalho, C.J.B. De & Couri, M.S. 1991. Muscidae, Fanniidae e Calliphoridae Silvio S. Nihei: Conceptualization, Data curation, (Diptera) do Projeto Maracá, Roraima, Brasil. Acta Amazonica, 21: Investigation, Methodology, Investigation, Project 35‑43. DOI administration, Resources, Supervision, Validation, Carvalho, C.J.B. De; Almeida, J.R. & Jesus, C.B. 1984. Dípteros sinantrópicos Visualization, Writing – original draft, and Writing – re- de Curitiba e arredores (Paraná, Brasil). I: Muscidae. Revista Brasileira de view & editing. Anderson T. 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APPENDIX: LIST OF EXAMINED MATERIAL

The species are listed in alphabetical order. Asterisk indicates new records to State of São Paulo. All material deposited at the Museu de Zoologia, Universidade de São Paulo, Brazil (MZSP).

Cyrtoneurina alifusca Couri, 1982: 1 male, 21.x‑19.xi.2010, Gudin & Nihei col.; 1 male, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 3 males, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 2 males, 21.i‑22.ii.2011, Gudin & Dios col.

Cyrtoneurina costalis (Walker, 1853): 1 male, 22.iii‑19.iv.2011, Moll & Nihei col.

Cyrtoneuropsis maculipennis Macquart, 1843: 1 female, 21.vii‑21.viii.2010, Moll & Nihei col.; 2 males, 1 female, 21.viii‑21.ix.2010, Gudin & Nihei col.; 3 females, 21.ix‑21.x.2010, Dios & Nihei col.; 1 male, 4 females, 21.x‑19.xi.2010, Gudin & Nihei col.; 1 male, 13 females, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 1 male, 8 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 3 males, 13 females, 21.i‑22.ii.2011, Gudin & Dios col.; 6 females, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.; 1 male, 3 females, 22.iii‑19.iv.2011, Moll & Nihei col.; 2 males, 6 females, 19.iv‑23.v.2011, Moll & Gudin col.; 3 females, 23.v‑20.vi.2011, Moll, Almeida & Tamakoshi col.; 2 males, 1 female, 20.vi‑23.vii.2011, Moll, Tamakoshi, Ribeiro col.

Cyrtoneuropsis polystigma (Wulp, 1896): 1 female, 21.vii‑21.viii.2010, Moll & Nihei col.; 2 males, 21.viii‑21.ix.2010, Gudin & Nihei col.; 4 males, 2 females, 21.ix‑21.x.2010, Dios & Nihei col.; 2 males, 4 females, 21.x‑19.xi.2010, Gudin & Nihei col.; 5 males, 4 females, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 4 males, 1 female, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 3 males, 4 females, 21.i‑22.ii.2011, Gudin & Dios col.

Dolichophaonia machadoi (Albuquerque, 1958)*: 4 females, 21.vii‑21.viii.2010, Moll & Nihei col.; 2 females, 21.viii‑21.ix.2010, Gudin & Nihei col.; 4 females, 21.x‑19.xi.2010, Gudin & Nihei col.; 2 females, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 1 female, 21.i‑22.ii.2011, Gudin & Dios col.; 1 female, 19.iv‑23.v.2011, Moll & Gudin col.; 1 female, 20.vi‑23.vii.2011, Moll, Tamakoshi, Ribeiro col.

Dolichophaonia simplex (Albuquerque, 1958)*: 1 female, 21.viii‑21.ix.2010, Gudin & Nihei col.

Helina aff. discreta (Wulp, 1896)*: 3 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 female, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.

Helina gigantea Albuquerque, 1956*: 2 females, 21.x‑19.xi.2010, Gudin & Nihei col.

Helina praecipua (Walker, 1853): 1 male, 23.v‑20.vi.2011, Moll, Almeida & Tamakoshi col.

Helina regobarrosi Albuquerque, 1958*: 1 male, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 1 female, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 female, 21.i‑22. ii.2011, Gudin & Dios col.

Morellia humeralis (Stein, 1919): 1 female, 21.i‑22.ii.2011, Gudin & Dios col.

Morellia nigricosta Hough, 1900: 3 females, 21.i‑22.ii.2011, Gudin & Dios col.

Mydaea plaumanni Snyder, 1941: 1 female, 21.ix‑21.x.2010, Dios & Nihei col.; 3 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 female, 19.iv‑23.v.2011, Moll & Gudin col.; 1 female, 23.v‑20.vi.2011, Moll, Almeida & Tamakoshi col.; 2 females, 20.vi‑23.vii.2011, Moll, Tamakoshi, Ribeiro col.

Myospila meditabunda (Fabricius, 1781): 1 female, 21.x‑19.xi.2010, Gudin & Nihei col.; 1 female, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.

Neomuscina anajeensis Pereira-Colavite & Carvalho 2012*: 1 female, 21.x‑19.xi.2010, Gudin & Nihei col.

Neomuscina atincticosta Snyder, 1949: 1 male, 21.xii.2010‑21.i.2011, Sato & Nihei col. Pap. Avulsos Zool., 2021; v.61: e20216168 Nihei, S.S. et al.: Muscidae from Paranapiacaba 12/12

Neomuscina inflexa (Stein, 1918): 2 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.

Neomuscina neosimilis Snyer, 1949: 1 female, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 1 female, 21.xii.2010‑21.i.2011, Sato & Nihei col.

Neomuscina schadei Snyder, 1949: 1 female, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.

Neomuscina transporta Snyder, 1949*: 1 female, 21.x‑19.xi.2010, Gudin & Nihei col.

Neurotrixa sulina Costacurta & Carvalho, 2005*: 1 female, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.

Phaonia advena Snyder, 1957*: 2 females, 21.vii‑21.viii.2010, Moll & Nihei col.; 2 males, 7 females, 21.viii‑21.ix.2010, Gudin & Nihei col.; 3 females, 21.ix‑21.x.2010, Dios & Nihei col.; 1 male, 5 females, 21.x‑19.xi.2010, Gudin & Nihei col.; 1 male, 13 females, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 5 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 male, 21.i‑22.ii.2011, Gudin & Dios col.; 4 females, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.; 2 females, 22.iii‑19.iv.2011, Moll & Nihei col.; 6 females, 19.iv‑23.v.2011, Moll & Gudin col.; 1 female, 23.v‑20.vi.2011, Moll, Almeida & Tamakoshi col.; 2 females, 20.vi‑23.vii.2011, Moll, Tamakoshi, Ribeiro col.

Phaonia annulata (Albuquerque, 1957)*: 1 female, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 male, 19.iv‑23.v.2011, Moll & Gudin col.

Phaonia grajauensis (Albuquerque, 1957): 2 female, 21.xii.2010‑21.i.2011, Sato & Nihei col.

Phaonia grandis (Couri, 1982)*: 1 male, 12 females, 21.vii‑21.viii.2010, Moll & Nihei col.; 3 males, 9 females, 21.viii‑21.ix.2010, Gudin & Nihei col.; 1 male, 6 fe‑ males, 21.ix‑21.x.2010, Dios & Nihei col.; 5 females, 21.x‑19.xi.2010, Gudin & Nihei col.; 11 females, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 1 male, 4 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 3 males, 13 females, 21.i‑22.ii.2011, Gudin & Dios col.; 1 male, 5 females, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.; 2 male, 5 females, 22.iii‑19.iv.2011, Moll & Nihei col.; 1 male, 11 females, 19.iv‑23.v.2011, Moll & Gudin col.; 8 females, 23.v‑20.vi.2011, Moll, Almeida & Tamakoshi col.; 2 males, 6 females, 20.vi‑23.vii.2011, Moll, Tamakoshi, Ribeiro col.

Phaonia lentiginosa Snyder 1957*: 2 males, 21.vii‑21.viii.2010, Moll & Nihei col.; 1 female, 21.viii‑21.ix.2010, Gudin & Nihei col.; 1 female, 21.ix‑21.x.2010, Dios & Nihei col.; 3 females, 21.x‑19.xi.2010, Gudin & Nihei col.; 1 male, 5 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 female, 21.i‑22.ii.2011, Gudin & Dios col.; 1 female, 22.iii‑19.iv.2011, Moll & Nihei col.; 2 females, 19.iv‑23.v.2011, Moll & Gudin col.

Phaonia shannoni (Carvalho & Pont, 1993): 1 female, 21.vii‑21.viii.2010, Moll & Nihei col.; 1 female, 21.viii‑21.ix.2010, Gudin & Nihei col.; 1 female, 21.i‑22. ii.2011, Gudin & Dios col.; 1 female, 20.vi‑23.vii.2011, Moll, Tamakoshi, Ribeiro col.

Phaonia similata (Albuquerque, 1957): 1 female, 21.viii‑21.ix.2010, Gudin & Nihei col.

Phaonia trispila (Bigot, 1885): 1 male, 1 female, 23.v‑20.vi.2011, Moll, Almeida & Tamakoshi col.

Philornis amazonenses Couri, 1983: 1 female, 21.viii‑21.ix.2010, Gudin & Nihei col.; 1 female, 21.i‑22.ii.2011, Gudin & Dios col.

Polietina minor (Albuquerque, 1956)*: 1 female, 21.i‑22.ii.2011, Gudin & Dios col.

Polietina orbitalis (Stein, 1904): 1 female, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.

Polietina prima (Couri & Machado, 1990): 2 females, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 female, 21.i‑22.ii.2011, Gudin & Dios col.; 2 females, 23.ii‑21. iii.2011, Moll, Nihei & Dias col.; 1 male, 22.iii‑19.iv.2011, Moll & Nihei col.; 1 female, 19.iv‑23.v.2011, Moll & Gudin col.

Polietina steini (Enderlein, 1927): 2 females, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.

Polietina univittata Couri & Carvalho, 1996: 1 female, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 2 females, 21.i‑22.ii.2011, Gudin & Dios col.; 1 female, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.

Pseudoptilolepis fulvapoda Snyder, 1949: 2 females, 21.viii‑21.ix.2010, Gudin & Nihei col.; 3 females, 21.ix‑21.x.2010, Dios & Nihei col.; 1 female, 21.x‑19. xi.2010, Gudin & Nihei col.; 1 female, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 1 male, 1 female, 21.xii.2010‑21.i.2011, Sato & Nihei col.; 1 female, 21.i‑22.ii.2011, Gudin & Dios col.; 1 female, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.; 1 female, 20.vi‑23.vii.2011, Moll, Tamakoshi, Ribeiro col.

Pseudoptilolepis nudapleura Snyder, 1949: 1 female, 21.xii.2010‑21.i.2011, Sato & Nihei col.

Psilochaeta pampiana (Shannon & Del Ponte, 1926): 1 male, 19.xi‑21.xii.2010, Gudin, Araya & Nihei col.; 1 male, 21.xii.2010‑21.i.2011, Sato & Nihei col.

Synthesiomyia nudiseta (Wulp, 1883): 1 female, 23.ii‑21.iii.2011, Moll, Nihei & Dias col.

Published with the nancial support of the "Programa de Apoio às Publicações Cientícas Periódicas da USP"